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Morphological characters and phylogenetic analysis reveal a new species of Sanghuangporus from China

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Abstract

Based on phylogenetic studies, the Inonotus linteus complex has been recently divided into two genera, Sanghuangporus and Tropicoporus. During investigations on the species diversity of the Inonotus linteus complex from China, a new species, Sanghuangporus quercicola sp. nov., is described based on morphological and molecular data. Morphologically, it is characterized by perennial, pileate basidiomata, a heterogeneous hyphal system with monomitic in context and dimitic in trama, and broadly subglobose to ovoid, thick-walled basidiospores measuring as 3–3.9 × 2.4–2.8 μm. Phylogenetically, the status of S. quercicola is strongly supported based on sequences of the nuclear internal transcribed spacer (ITS) regions.

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... Besides morphological and ecological (host preference) characters, the ITS barcoding region provides the most powerful tool for differentiating species of the genus. For example, more than half of the known species of Sanghuangporus were discovered with the aid of the ITS region alone (Wu et al. 2012a(Wu et al. , 2019bTian et al. 2013;Ghobad-Nejhad 2015;Tomšovský 2015;Zhu et al. 2017). Moreover, the reliability of the ITS region for species differentiation in the genus has been substantiated by a multilocus-based phylogenetic analysis (Zhu et al. 2019). ...
... Sanghuangporus toxicodendri was recently described from specimens collected from Toxicodendron sp. in Hubei, central China (Wu et al. 2019b) and resembles S. quercicola, another species originally described from central China (Zhu et al. 2017). However, in the publication introducing S. toxicodendri (Wu et al. 2019b) the separation from S. quercicola was not well-supported phylogenetically. ...
... However, in the publication introducing S. toxicodendri (Wu et al. 2019b) the separation from S. quercicola was not well-supported phylogenetically. Moreover, the morphological differences between these two species are slight (such as for basidiospore length) or involve variable characters that do not have taxonomic signal (such as the surface color of the pileal margin) (Zhu et al. 2017;Wu et al. 2019b). In the current phylogenetic analyses, the six specimens of S. toxicodendri, three specimens of S. quercicola and four additional collections merged in a fully supported clade (Additional file 1: Tree S1, Additional file 2: Tree S2, Fig. 2). ...
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“Sanghuang” refers to a group of important traditionally-used medicinal mushrooms belonging to the genus Sanghuangporus . In practice, species of Sanghuangporus referred to in medicinal studies and industry are now differentiated mainly by a BLAST search of GenBank with the ITS barcoding region as a query. However, inappropriately labeled ITS sequences of “Sanghuang” in GenBank restrict accurate species identification and, to some extent, the utilization of these species as medicinal resources. We examined all available 271 ITS sequences related to “Sanghuang” in GenBank including 31 newly submitted sequences from this study. Of these sequences, more than half were mislabeled so we have now corrected the corresponding species names. The mislabeled sequences mainly came from strains utilized by non-taxonomists. Based on the analyses of ITS sequences submitted by taxonomists as well as morphological characters, we separate the newly described Sanghuangporus subbaumii from S. baumii and treat S. toxicodendri as a later synonym of S. quercicola . Fourteen species of Sanghuangporus are accepted, with intraspecific distances up to 1.30% (except in S. vaninii , S. weirianus and S. zonatus ) and interspecific distances above 1.30% (except between S. alpinus and S. lonicerinus , and S. baumii and S. subbaumii ). To stabilize the concept of these 14 species of Sanghuangporus , their taxonomic information and reliable ITS reference sequences are provided. Moreover, ten potential diagnostic sequences are provided for Hyperbranched Rolling Circle Amplification to rapidly confirm three common commercial species, viz. S. baumii , S. sanghuang , and S. vaninii . Our results provide a practical method for ITS barcoding-based species identification of Sanghuangporus and will promote medicinal studies and commercial development from taxonomically correct material.
... Sanghuangporus sanghuang (Sheng H. Wu et al.) Sheng H. Wu et al., the generic type, was detected by Wu et al. (2012) as the genuine Sanghuang species growing exclusively on Morus in the wild. Before this study, 13 species of Sanghuangporus were known (Ghobad-Nejhad 2015;Tomsovsky 2015;Zhou et al. 2016;Zhu et al. 2017). In this study, we present a new species of Sanghuangporus sp. ...
... The sequences were then submitted to Genbank. Zhu et al. (2017) conducted ITS-based phylogenetic analysis for all previously known 13 species of Sanghuangporus. The ingroup strains of the Sanghuangporus spp. ...
... ( Zhu et al.'s (2019) phylogenetic study showed the monophyly of the genus Sanghuangporus spp., and the result coincides with the present study (Fig. 1). The genus Sanghuangporus comprises 14 species (Ghobad-Nejhad 2015;Tomsovsky 2015;Zhou et al. 2016;Zhu et al. 2017), after including S. toxicodendri presented here. It is not easy to identify some species of Sanghuangporus spp., as there are not that many good morphological characteristics to separate them. ...
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Sanghuangporus toxicodendri (Hymenochaetales) is described as new based on collections made from Shennongjia Forestry District, Hubei Province, China. All studied basidiocarps grew on living trunks of Toxicodendron sp. This new species is characterized by having perennial, effused-reflexed to pileate basidiocarps; pore surface brownish yellow or yellowish brown, pores 7-9 per mm; context 1-5 mm thick or almost invisible; setae ventricose, dark brown, 26-42 × 7-10 μm; basidia 4-sterigmate or occasionally 2-sterigmate; basidiospores broadly ellipsoid, smooth, brownish yellow, slightly thick-walled, mostly 3.5-4 × 2.8-3 μm. Maximum likelihood and Bayesian inference phylogenies inferred from internal transcribed spacer (ITS) region of rDNA indicated that Sanghuangporus spp. formed a monophyletic clade and resolved as a sister to Tropicoporus spp., and six strains of S. toxicodendri formed a monophyletic group which is sister to S. quercicola. An identification key to known species of Sanghuangporus is provided.
... Species in this genus form parasitic relationships with Alnus, Juglans, Lonicera, Morus, Populus, Quercus, Syringa, and Weigela (Dai, 2010;Wu et al., 2012). In the past few years, several studies have examined the phylogenetic relationships within Sanghuangporus (Dai, 2010;Wu et al., 2012;Vlasák et al., 2013;Tian et al., 2013;Ghobad-Nejhad, 2015;Tomsovsky, 2015;Zhou et al., 2016;Zhu et al., 2017). Thirteen species are currently accepted in the genus: S. alpinus (Y.C. ...
... Thirteen species are currently accepted in the genus: S. alpinus (Y.C. Dai (Dai, 2010;Wu et al., 2012;Tian et al., 2013;Vlasák et al., 2013;Ghobad-Nejhad, 2015;Tomsovsky, 2015;Zhou et al., 2016;Zhu et al., 2017). ...
... (3) basidiospores ovoid to broadly ellipsoid Tomsovsky, 2015;Zhu et al., 2017). Nevertheless, S. quercicola grows on Quercus, but S. pilatii and S. vaninii mostly grow on Populus Tomsovsky, 2015;Zhu et al., 2017). ...
Article
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“Sanghuang” is a popular fungus used as a Chinese traditional medicine. In fact, it represents a group of fungi belonging to the genus Sanghuangporus, but little is known about its origin and biogeography. The aim of this study was to characterize the molecular relationships, origin and biogeographical distribution of Sanghuangporus. The multi-locus phylogenetic analyses were used to infer the phylogenetic relationships. In addition, based on Bayesian evolutionary analysis using sequences from the internal transcribed spacer (ITS), nuclear large subunit rDNA (nLSU), translation elongation factor 1-α (EF1-α), and the largest and second largest subunits of RNA polymerase II (RPB1 and RPB2), we used a fungus fossil-based approach to gain insight into the divergence time of species in Sanghuangporus. The molecular phylogeny strongly supports the monophyly of Sanghuangporus (MP = 100%, ML = 100%, and BPP = 1.00), and 13 species are recognized in this genus. The Bayesian uncorrelated lognormal relaxed molecular clock using BEAST and reconstructed ancestral areas indicate that the maximum crown age of Sanghuangporus is approximately 30.85 million years. East Asia is the likely ancestral area (38%). Dispersal and differentiation to other continents then occurred during the late Middle Miocene and Pliocene. The ancestor of Sanghuangporus probably originated in palaeotropical Northeast Asia and covered Northeast Asia and East Africa during the Oligocene-Miocene, hosted by plants that expanded via the “Gomphotherium Landbridge.” Six kinds of dispersal routes are proposed, including intercontinental dispersal events of three clades between Northeast Asia and East Africa, between East Asia and North America, and between Northeast Asia and Europe.
... In this study, we present a new species of Sanghuangporus from tropical South Taiwan, that grows on the trunk of living Vitex negundo. The genus Sanghuangporus accommodates 14 species (Ghobad-Nejhad 2015;Tomsovsky 2015;Zhou et al. 2016;Zhu et al. 2017;Zhu et al. 2019) including the new species, and a key to these species is provided. ...
... (Wu et al. 2019) while the latter on Quercus sp. (Zhu et al. 2017). In addition to different host tree genus (also different family), two morphological features can separate these two species, according to Wu et al. (2019). ...
Article
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Sanghuangporus vitexicola (Hymenochaetales) is described as a new species based on collections made from Pingtung County, tropical South Taiwan. All studied basidiocarps grew on living trunks of Vitex negundo. This new species is characterized by having perennial, pileate basidiocarps; pore surface yellowish brown, pores 6–8 per mm; context 0.7–1.5 cm thick; setae ventricose to subulate, dark brown, 17–30 × 5–8 μm; basidiospores broadly ellipsoid, brownish, smooth, 4.2–4.8 × 3.2–3.7 μm, with 0.2–0.5 μm thick walls. Maximum likelihood and Bayesian inference phylogenies inferred from the internal transcribed spacer (ITS) region of rDNA indicated that six strains of Sanghuangporus viexicola formed a monophyletic group which is sister to S. zonatus. An identification key to known species of Sanghuangporus is provided.
... The characteristics of the three different developmental stages (20-day-old mycelia and fruiting bodies 1 and 3 years in age) were evaluated (Fig. 1a). S. vaninii Kangneng fruiting bodies had a brown or dark grayish to black pileal surface with a distinct yellow margin, consistent with the characteristics of S. vaninii previously reported (Zhu et al. 2017). A total of 9956 genes were detected in the sequenced samples, accounting for approximately 88.03% of the proteinencoding genes annotated (Table S1). ...
Article
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The medicinal fungus Sanghuang produces diverse bioactive compounds and is widely used in Asian countries. However, little is known about the genes and pathways involved in the biosynthesis of these active compounds. Based on our previous study providing Sanghuangporus vaninii genomic information, the transcriptomes of MY (mycelium), OY (1-year-old fruiting bodies), and TY (3-year-old fruiting bodies) were determined in this study. A significant number of genes (4774) were up- or downregulated between mycelium and fruiting bodies, but only 1422 differentially expressed genes were detected between OY and TY. 138 genes encoding P450s were identified in the fungal genome and grouped into 25 P450 families; more than 64% (88) of the genes were significantly differentially expressed between the mycelium and fruiting body, suggesting that these P450s are involved in fungal sexual development. Importantly, the expression of genes involved in bioactive compound (triterpenoids, polysaccharides, and flavonoids) biosynthesis in asexual (cultured with solid and liquid media) and sexual stages was explored and combined with transcriptome and quantitative PCR analyses. More genes involved in the biosynthesis of bioactive compounds were expressed more highly in mycelium than in fruiting bodies under liquid medium culture compared with solid medium culture, which was consistent with the yields of different bioactive compounds, suggesting that liquid fermentation of S. vaninii Kangneng can be used to obtain these bioactive compounds. A comprehensive understanding of the genomic information of S. vaninii will facilitate its potential use in pharmacological and industrial applications.
... This taxonomic system is widely acknowledged and soon four new members were added to Sanghuangporus. [22][23][24][25] To date, 14 species distributed around the world are accepted in Sanghuangporus and are considered to be members of 'Sanghuang' (Table 1, Fig. 1). Noteworthily, members of Tropicoporus, including T. linteus (Berk. ...
Article
‘Sanghuang’ is one of the most important groups of medicinal macrofungi and has been used in traditional Chinese medicine for the past two centuries. However, knowledge about ‘Sanghuang’ is distributed in journals of different disciplines, which has meant it is not well known. To facilitate the benefit of ‘Sanghuang’ to human health, here we summarize its current status as a group of medicinal macrofungi. ‘Sanghuang’ consists of all members of the genus Sanghuangporus, which now accommodates 14 described species distributed around the world. The powders or chips of basidiocarps as supplements are the main method of consumption. In the case that wild basidiocarps of ‘Sanghuang’ are not sufficient, cultivation is supposed to provide a solution to promote the long-term utilization of ‘Sanghuang’ basidiocarps. ‘Sanghuang’ mainly possesses antitumor, antioxidant, anti-inflammatory, immunomodulatory, antidiabetic, hepatoprotective, and other medicinal properties. Polysaccharides, polyphenols, pyrones, and terpenes are the main metabolites responsible for these medicinal functions. Finally, four directions are proposed to improve the sustainable utilization of ‘Sanghuang’ resources, viz. accurate identification and standard deposition, documentation of species diversity, multi-omics analyses, and a cover-all database. Hopefully, this review will draw more attention to the utilization of ‘Sanghuang’ as medicinal mushrooms in a large scale of industry.
... was determined to be sanghuang (Dai and Xu, 1998), and nearly all morphologically similar materials were named as I. baumii. However, subsequent studies on systematics segregated several cryptic species from I. baumii (Wu et al., 2012;Zhu et al., 2017). Therefore, medicinal functions once considered to be possessed by P. linteus and I. baumii could be an inappropriate summarization from several cryptic species of genuine sanghuang under the incorrect names as indicated in Han et al. (2016) or of some other species. ...
Article
Recently, it has been indicated that systematics is of no value to the traditional Chinese medicinal studies and industry of macrofungi. As a fungal taxonomist, I use three genera of the most famous medicinal macrofungi, viz. Sanghuangporus, Ganoderma and Ophiocordyceps, as examples to rebut this opinion. Generally, the well-established systematics are helpful in using correct Latin names for fungal species in medicinal studies; furthermore, the resulting medicinal functions can be exactly matched to a particular species; finally, the particular species can be applied for permissions of utilization in traditional Chinese medicinal industry. Therefore, systematics is crucial for unlocking the potential of macrofungi as valuable resources to be developed in the traditional Chinese medicinal studies and industry, and should continue to be of interest to not just fungal taxonomists but also biochemists, pharmacists and businessmen.
... These cryptic species are included here and their functions are labeled following the phylogenetically closely related species. (Wu et al. 2012b;Tian et al. 2013;Zhou et al. 2016a;Zhu et al. 2017b . In the case that 173 species are newly included, but the number of Table 1). ...
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The huge land areas in China provide highly diverse habitats for macrofungi. Of these macrofungi, many are directly related to people’s daily life and have been utilized by ancient Chinese for at least 6800 years. In this study, we evaluate the current known resource diversity of Chinese macrofungi. A total of 1662 taxa are summarized, and all species names and their authorities have been checked and corrected according to authentic mycological databases. Among the 1662 taxa, 1020, 692, and 480 are considered to be edible, medicinal and poisonous mushrooms, respectively. A few of edible macrofungi in China are commonly used for commercial production. All known medicinal functions are labeled for medicinal species. The most common medicinal functions possessed by Chinese macrofungi are antitumor or anticancer, followed by antioxidant and antimicrobial. A total of 277 Chinese macrofungi are edible simultaneously with certain medicinal functions and without known toxicity. These species could be treated as “Gold Mushrooms”. Contrarily, 193 edible and/or medicinal species are also recognized as poisonous mushrooms. To avoid poisoning caused by these species, ingestion either in a proper way or in small amounts is important. However, the mycotoxins metabolized by these poisonous species could be a huge wealth of natural products yet to be explored. How to utilize these Chinese macrofungal resources is a critical to benefit humans worldwide.
... It originally included S. alpinus, S. baumii, S. lonicericola, S. lonicerinus, S. microcystideus, S. sanghuang, S. vaninii, S. weigelae, S. weirianus and S. zonatus (Zhou et al., 2015). Three more species have been added later, viz. S. ligneus, S. pilatii and S. quercicola (Ghobad-Nejhad, 2015;Tomsovsky, 2015;Zhu et al., 2017). The Asian species have been referred to in the literature as medicinal mushrooms. ...
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Taxonomy and phylogeny of poroid Hymenochaetaceae based on the most comprehensive phylogenetic analyses are presented. A phylogeny based on a combined dataset of ITS and nLSU sequences for accepted genera of Hymenochaetaceae was analyzed and multigene phylogenies for most species of ten large genera including Clotricia , Fomitiporella , Fomitiporia , Fulvifomes , Fuscoporia , Inonotus , Phylloporia , Porodaedalea , Sanghuangporus and Tropicoporus , were carried out. Based on samples from 37 countries of five continents, seven new genera, Meganotus, Neophellinus, Nothonotus, Pachynotus, Perenninotus, Pseudophylloporia and Rigidonotus, are introduced, 37 new species, Coltricia tibetica , Fomitiporella crassa , F. queenslandica , Fomitiporia eucalypti, F. gatesii , F. ovoidospora , Fulvifomes azonatus, F. caligoporus , F. costaricense , F. floridanus , F. jouzaii , F. nakasoneae , F. subindicus , Fuscoporia sinuosa , F. submurina , Inonotus subradiatus , I. vietnamensis , Neomensularia castanopsidis , Pachynotus punctatus , Phellinus cuspidatus , P. subellipsoideus , Phylloporia minutissima , P. tabernaemontanae , Porodaedalea occidentiamericana , P. orientoamericana , P. qilianensis , P. schrenkianae , Pseudophylloporia australiana , Sanghuangporus australianus , S. lagerstroemiae , Tropicoporus angustisulcatus , T. hainanicus , T. lineatus , T. minus , T. ravidus , T. substratificans and T. tenuis, are described, and 108 new combinations are proposed. In addition, one illegitimate name and two invalid names are renamed. The taxonomic relevance and limits of the new taxa are discussed. Photos and illustrations for 37 new species are presented, and a full description for each new species is given. Eventually, this study recognizes 672 species in 34 genera and provides a modern treatment of the poroid Hymenochaetaceae in the world. A key to the accepted poroid genera of Hymenochaetaceae is provided, and identification keys to the accepted species of 32 poroid genera worldwide are given. A synopsis description of each species is included in these keys.
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A legendary and very valuable medicinal fungus first known in China 2000 years ago is recognized in this study as a new species. The sanghuang mushroom (sanghuang) is a popular medicinal polypore used throughout China, Japan, and Korea. While its medicinal properties were recognized in an early Tang Dynasty herbal written in the 7th century, modern scientific research has only recently confirmed its highly effective antitumor properties. Although Japanese and Korean mycologists have adopted Phellinus linteus or P. baumii as the scientific name for sanghuang that belongs to the Inonotus baumii-I. linteus group in the Hymenochaetales (Basidiomycota), its species identity has not been satisfactorily answered. This study delimits the species of the I. baumii-I. linteus group including sanghuang, based on an analysis of morphological characteristics and nrDNA ITS sequences. Both morphological and molecular features were useful in separating different fungal species in this monophyletic group that are generally specialized with their host tree species. The true sanghuang is a new and previously undescribed species that grows solely on Morus in China, Japan, Korea, and Taiwan. It is now rare and endangered in the wild. Six related species of the I. baumii-I. linteus group distributed in Asia have evolved to specific host tree species: I. baumii on Syringa, I. lonicericola on Lonicera, I. lonicerinus comb. nov. on Lonicera, I. sanghuang on Morus, I. vaninii on Populus, and I. weigelae sp. nov. on Weigela; a key is provided to ease the determination of these taxa. Up to 1 more results found for "Species clarification for the medicinally valuable ‘sanghuang’ mushroom"
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The recently-developed statistical method known as the "bootstrap" can be used to place confidence intervals on phylogenies. It involves resampling points from one's own data, with replacement, to create a series of bootstrap samples of the same size as the original data. Each of these is analyzed, and the variation among the resulting estimates taken to indicate the size of the error involved in making estimates from the original data, In the case of phylogenies, it is argued that the proper method of resampling is to keep all of the original species while sampling characters with replacement, under the assumption that the characters have been independently drawn by the systematist and have evolved independently. Majority-rule consensus trees can be used to construct a phylogeny showing all of the inferred monophyletic groups that occurred in a majority of the bootstrap samples. If a group shows up 95% of the time or more, the evidence for it is taken to be statistically significant. Existing computer programs can be used to analyze different bootstrap samples by using weights on the characters, the weight of a character being how many times it was drawn in bootstrap sampling. When all characters are perfectly compatible, as envisioned by Hennig, bootstrap sampling becomes unnecessary; the bootstrap method would show significant evidence for a group if it is defined by three or more characters.
Dissepiments distinctly thin to slightly thick (<1/4 diameter of pores); distribution in Eastern Asia
  • ................... . Asia
Dissepiments distinctly thick; distribution in Western Asia............................................................................................... S. ligneous 10. Dissepiments distinctly thin to slightly thick (<1/4 diameter of pores); distribution in Eastern Asia...................................S. baumii 11. Basidiospores basically subglobose; restricted to Juglans...............................................................................................S. weirianus 11. Basidiospores broadly ellipsoid; restricted to Morus or Populus.....................................................................................................12 12.
Basidiospores 4-4.9 × 3.1-3.9 μm; restricted to Morus
  • . . . Populus
Basidiospores 3.8-4.4 × 2.8-3.7 μm; restricted to Populus..................................................................................................S. vaninii 12. Basidiospores 4-4.9 × 3.1-3.9 μm; restricted to Morus................................................................................................. S. sanghuang References Chen, J.J., Cui, B.K. & Dai, Y.C. (2016) Global diversity and molecular systematics of Wrightoporia s. l. (Russulales, Basidiomycota).
Farvekort. The Danish mycological society's colour-chart. Foreningentil Svampekundskabens Fremme, Greve
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Petersen, J.H. (1996) Farvekort. The Danish mycological society's colour-chart. Foreningentil Svampekundskabens Fremme, Greve. pp. 1-6.
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Song, J., Chen, J.J., Wang, M., Chen, Y.Y. & Cui, B.K. (2016) Phylogeny and biogeography of the remarkable genus Bondarzewia (Basidiomycota, Russulales). Scientific Reports 6: 34568.
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Zhou, L.W., Vlasák, J., Decock, C., Asefa, A., Stelid, J., Abate, D., Wu, S.H. & Dai, Y.C. (2016) Global diversity and taxonomy of the Inonotus linteus complex (Hymenochaetales, Basidiomycota): Sanghuangporus gen. nov., Tropicoporus excentrodendri and T. guanacastensis gen. et spp. nov., and 17 new combinations. Fungal Diversity 77: 335-347.